Genes & Genomics

, Volume 41, Issue 2, pp 223–231 | Cite as

Study of QTLs linked to awn length and their relationships with chloroplasts under control and saline environments in bread wheat

  • Bahram MasoudiEmail author
  • Mohsen Mardi
  • Eslam Majidi Hervan
  • Mohammad Reza Bihamta
  • Mohammad Reza Naghavi
  • Babak Nakhoda
  • Behnam Bakhshi
  • Mehrzad Ahmadi
  • Mohammad Taghi Tabatabaei
  • Mohamad Hossein Dehghani Firouzabadi
Research Article



Some studies in wheat showed that awns may have a useful effect on yield, especially under drought stress. Up to this time few researches has identified the awn length QTLs with different effect in salinity stress.


The primary objective of this study was to examine the additive (a) and the epistatic (aa) QTLs involve in wheat awns length in control and saline environments.


A F7 RIL population consisting of 319 sister lines, derived from a cross between wheat cultivars Roshan and Falat (seri82), and the two parents were grown in two environments (control and Saline) based on an alpha lattice design with two replications in each environment. At flowering, awn length was measured for each line. For QTL analysis, the linkage map of the ‘‘Roshan × Falat’’ population was used, which included 748 markers including 719 DArT, 29 simple sequenced repeats (SSRs). Additive and pleiotropic QTLs were identified. In order to reveal the relationship between the identified QTL for awns length and the role of the gene or genes that it expresses, the awns length locus location and characteristics of its related CDS, gene, UTRs, ORF, exons and Introns were studied using ensemble plant ( Furthermore, the promoter analysis has been done using NSITE-PL.


We identified 6 additive QTLs for awn length by QTL Cartographer program using single-environment phenotypical values. Also, we detected three additive and two epistatic QTLs for awn length by the QTLNetwork program using multi-environment phenotypical values. Our results showed that none of the additive and epistatic QTLs had interactions with environment. One of the additive QTLs located on chromosome 4A was co-located with QTLs for number of sterile spikelet per spike in both environment and number of seed per spike in control environment.


Studies of the locus linked to the awns length QTL revealed the role of awn and its chloroplasts in grain filing during abiotic stress could be enhanced by over expression of some genes like GTP-Binding proteins which are enriched in chloroplasts encoded by genes included wPt-5730 locus.


Awns length Quantitative trait loci Epistasis effects Salt tolerance Wheat 



This research was supported by Agricultural Biotechnology Research Institute of Iran (ABRII).


  1. Abebe T, Melmaiee K, Berg V, Wise RP (2010) Drought response in the spikes of barley: gene expression in the lemma, palea, awn, and seed. Funct Integr Gen 10:191–205CrossRefGoogle Scholar
  2. Araus J, Brown H, Febrero A, Bort J, Serret M (1993) Ear photosynthesis, carbon isotope discrimination and the contribution of respiratory CO2 to differences in grain mass in durum wheat. Plant Cell Environ 16:383–392CrossRefGoogle Scholar
  3. Ban T, Suenaga K (2000) Genetic analysis of resistance to Fusarium head blight caused by Fusarium graminearum in Chinese wheat cultivar Sumai 3 and the Japanese cultivar Saikai 165. Euphytica 113:87–99CrossRefGoogle Scholar
  4. Biscoe P, Littleton E, Scott R (1973) Stomatal control of gas exchange in barley awns. Ann Appl Biol 75:285–297CrossRefGoogle Scholar
  5. Blum A (1985) Photosynthesis and transpiration in leaves and ears of wheat and barley varieties. J Exp Bot 36:432–440CrossRefGoogle Scholar
  6. Bort J, Brown RH, Araus JL (1996) Refixation of respiratory CO2 in the ears of C3 cereals. J Exp Bot 47:1567–1575CrossRefGoogle Scholar
  7. Carlborg Ö, Jacobsson L, Åhgren P, Siegel P, Andersson L (2006) Epistasis and the release of genetic variation during long-term selection. Nat Genet 38:418–420CrossRefGoogle Scholar
  8. Duffus C, Cochrane M (1993) Formation of the barley grain: morphology, physiology, and biochemistry. In: MacGregor AW, Bhatty RS (eds) Barley: Chemistry and Technology. American Association of Cereal Chemists, St Paul, Minn, pp 31–72Google Scholar
  9. Evans L, Rawson HM (1970) Photosynthesis and respiration by the flag leaf and components of the ear during grain development in wheat. Australian J Biol Sci 23:245–254CrossRefGoogle Scholar
  10. Evans L, Bingham J, Jackson P, SUTHERLAND J (1972) Effect of awns and drought on the supply of photosynthate and its distribution within wheat ears. Ann Appl Biol 70:67–76CrossRefGoogle Scholar
  11. Gale M, Atkinson M, Chinoy C, Harcourt R, Jia J, Li Q, Devos K (1995) Genetic maps of hexaploid wheat. In: Proc 8th Int Wheat Genet Symp. China Agricultural Scientech Press, Beijing, pp 29–40Google Scholar
  12. Grundbacher F (1963) The physiological function of the cereal awn. Bot Rev 29:366–381CrossRefGoogle Scholar
  13. Guo Z, Schnurbusch T (2016) Costs and benefits of awns. J Exp Bot 67:2533–2535CrossRefGoogle Scholar
  14. Guo Z, Chen D, Schnurbusch T (2015) Variance components, heritability and correlation analysis of anther and ovary size during the floral development of bread wheat. J Exp Bot 66(11):3099–111CrossRefGoogle Scholar
  15. Hein JA, Sherrard ME, Manfredi KP, Abebe T (2016) The fifth leaf and spike organs of barley (Hordeum vulgare L.) display different physiological and metabolic responses to drought stress. BMC Plant Biol 16:248CrossRefGoogle Scholar
  16. Jia S, Lv J, Jiang S, Liang T, Liu C, Jing Z (2015) Response of wheat ear photosynthesis and photosynthate carbon distribution to water deficit. Photosynthetica 53:95–109CrossRefGoogle Scholar
  17. Jiang C, Zeng Z-B (1995) Multiple trait analysis of genetic mapping for quantitative trait loci. Genetics 140:1111–1127Google Scholar
  18. Koyama ML, Levesley A, Koebner RM, Flowers TJ, Yeo AR (2001) Quantitative trait loci for component physiological traits determining salt tolerance in rice. Plant Physiol 125:406–422CrossRefGoogle Scholar
  19. Li X-f, Hong-Gang BD W (2010) Awn anatomy of common wheat (Triticum aestivum L.) and its relatives. Caryologia 63:391–397CrossRefGoogle Scholar
  20. Li Z, Yu S, Lafitte H, Huang N, Courtois B, Hittalmani S, Vijayakumar C, Liu G, Wang G, Shashidhar H (2003) QTL × environment interactions in rice. I. Heading date and plant height. Theor Appl Genet 108:141–153CrossRefGoogle Scholar
  21. Liu Y-G, Tsunewaki K (1991) Restriction fragment length polymorphism (RFLP) analysis in wheat. II. Linkage maps of the RFLP sites in common wheat. Idengaku zasshi 66:617–633Google Scholar
  22. Ma L, Zhou E, Huo N, Zhou R, Wang G, Jia J (2007) Genetic analysis of salt tolerance in a recombinant inbred population of wheat (Triticum aestivum L.). Euphytica 153:109–117CrossRefGoogle Scholar
  23. Malmberg RL, Mauricio R (2005) QTL-based evidence for the role of epistasis in evolution. Genet Res 86:89–96CrossRefGoogle Scholar
  24. Malmberg RL, Held S, Waits A, Mauricio R (2005) Epistasis for fitness-related quantitative traits in Arabidopsis thaliana grown in the field and in the greenhouse. Genetics 171:2013–2027CrossRefGoogle Scholar
  25. Manneh B, Stam P, Struik PC, Bruce-Oliver S, Van Eeuwijk FA (2007) QTL-based analysis of genotype-by-environment interaction for grain yield of rice in stress and non-stress environments. Euphytica 156:213–226CrossRefGoogle Scholar
  26. Maydup ML, Antonietta M, Guiamet J, Graciano C, López JR, Tambussi EA (2010) The contribution of ear photosynthesis to grain filling in bread wheat (Triticum aestivum L.). Field Crops Res 119:48–58CrossRefGoogle Scholar
  27. Maydup M, Antonietta M, Graciano C, Guiamet J, Tambussi E (2014) The contribution of the awns of bread wheat (Triticum aestivum L.) to grain filling: responses to water deficit and the effects of awns on ear temperature and hydraulic conductance. Field crops research 167:102–111CrossRefGoogle Scholar
  28. McDonough WT, Gauch HG (1959) Contribution of the awns to the development of the kernels of bearded wheat. Maryland Ag Exp Sta Bull A103:1–16Google Scholar
  29. McIntosh R, Hart G, Devos K, Gale M, Rogers W (1998) Catalogue of gene symbols for wheat. In: Proc 9th Int Wheat Genet Symp Saskatoon, vol 5, pp 235Google Scholar
  30. Merah O, Deléens É, Teulat B, Monneveux P (2001) Productivity and carbon isotope discrimination in durum wheat organs under a Mediterranean climate. Comptes Rendus de lAcadémie des Sciences-Series III-Sciences de la Vie 324:51–57CrossRefGoogle Scholar
  31. Miller E, Gauch H, Gries G (1944) A study of the morphological nature and physiological function of the awns in winter wheat. Kansas Agric Exp Stn Tech Bull 57:1–82Google Scholar
  32. Morgan J (1980) Osmotic adjustment in the spikelets and leaves of wheat. J Exp Bot 31:655–665CrossRefGoogle Scholar
  33. Motzo R, Giunta F (2002) Awnedness affects grain yield and kernel weight in near-isogenic lines of durum wheat. Crop Pasture Sci 53:1285–1293CrossRefGoogle Scholar
  34. Nelson JC, Deynze AEV, Sorrells ME, Autrique E, Lu YH, Merlino M, Atkinson M, Leroy P (1995a) Molecular mapping of wheat. Homoeologous group 2. Genome 38:516–524CrossRefGoogle Scholar
  35. Nelson JC, Deynze AEV, Sorrells ME, Autrique E, Lu YH, Negre S, Bernard M, Leroy P (1995b) Molecular mapping of wheat. Homoeologous group 3. Genome 38:525–533CrossRefGoogle Scholar
  36. Nelson JC, Sorrells ME, Van-Deynze A, Lu YH, Atkinson M, Bernard M, Leroy P, Faris JD, Anderson JA (1995c) Molecular mapping of wheat: major genes and rearrangements in homoeologous groups 4, 5, and 7. Genetics 141:721Google Scholar
  37. Paterson AH (1995) Molecular dissection of quantitative traits: progress and prospects. Genome Res 5:321–333CrossRefGoogle Scholar
  38. Poustini K, Siosemardeh A (2004) Ion distribution in wheat cultivars in response to salinity stress. Field Crops Res 85:125–133CrossRefGoogle Scholar
  39. Rao P (1981) Telocentric mapping of the awn inhibitor gene Hd on chromosome 4B of common wheat. Cereal Res Commun 9(4):335–337Google Scholar
  40. Rebetzke G, Bonnett D, Reynolds M (2016) Awns reduce grain number to increase grain size and harvestable yield in irrigated and rainfed spring wheat. J Exp Bot 67:2573–2586CrossRefGoogle Scholar
  41. Sears ER (1954) The aneuploids of common wheat, vol 572. University of Missouri, College of Agriculture, Agricultural Experiment Station, Columbia, Mo, pp 1–58Google Scholar
  42. Sears E (1966) Chromosome mapping with the aid of telocentrics. In: Proc 2nd Int wheat genet symp. hereditas suppl, pp 370–381Google Scholar
  43. Shen X, Zhang T, Guo W, Zhu X, Zhang X (2006) Mapping fiber and yield QTLs with main, epistatic, and QTL × environment interaction effects in recombinant inbred lines of upland cotton. Crop Sci 46:61–66CrossRefGoogle Scholar
  44. Snijders C (1990) Fusarium head blight and mycotoxin contamination of wheat, a review. Neth J Plant Pathol 96:187–198CrossRefGoogle Scholar
  45. Solovyev VV, Shahmuradov IA, Salamov AA (2010) Identification of promoter regions and regulatory sites. In: Computational biology of transcription factor binding. Springer, Berlin, pp 57–83Google Scholar
  46. Sourdille P, Cadalen T, Gay G, Gill B, Bernard M (2002) Molecular and physical mapping of genes affecting awning in wheat. Plant Breed 121:320–324CrossRefGoogle Scholar
  47. Steiner B, Lemmens M, Griesser M, Scholz U, Schondelmaier J, Buerstmayr H (2004) Molecular mapping of resistance to Fusarium head blight in the spring wheat cultivar Frontana. Theor Appl Genet 109:215–224CrossRefGoogle Scholar
  48. Suwastika IN, Denawa M, Hata A, Ohniwa RL, Takeyasu K, Shiina T (2008) Localization of Obg-Hflx and TrmE-Era super family small GTPases in various organelles in plant cells. In: Photosynthesis. Energy from the Sun. Springer, pp 1137–1140Google Scholar
  49. Tamburic-Ilincic L, Schaafsma A, Falk D (2007) Indirect selection for lower deoxynivalenol (DON) content in grain in a winter wheat population. Can J Plant Sci 87:931–936CrossRefGoogle Scholar
  50. Tanksley SD (1993) Mapping polygenes. Annu Rev Genet 27:205–233CrossRefGoogle Scholar
  51. TORNE GN (1963) Varietal differences in photosynthesis of ears and leaves of barley. Ann Bot 27:155–174CrossRefGoogle Scholar
  52. Veldboom L, Lee M, Woodman W (1994) Molecular marker-facilitated studies in an elite maize population: I. Linkage analysis and determination of QTL for morphological traits. Theor Appl Genet 88:7–16CrossRefGoogle Scholar
  53. Vervelde G (1953) The agricultural value of awns in cereals. Neth J Agric Sci 1(2):2–10Google Scholar
  54. Watkins A, Ellerton S (1940) Variation and genetics of the awn in Triticum. J Genet 40:243–270CrossRefGoogle Scholar
  55. Weyhrich RA, Carver BF, Martin BC (1995) Photosynthesis and water-use efficiency of awned and awnletted near-isogenic lines of hard winter wheat. Crop Sci 35:172–176CrossRefGoogle Scholar
  56. Würschum T, Maurer HP, Schulz B, Möhring J, Reif JC (2011) Genome-wide association mapping reveals epistasis and genetic interaction networks in sugar beet. Theor Appl Genet 123:109–118CrossRefGoogle Scholar
  57. Xie Q, Mayes S, Sparkes DL (2015) Carpel size, grain filling, and morphology determine individual grain weight in wheat. J Exp Bot 66(21):6715–6730CrossRefGoogle Scholar
  58. Xu Y-F, An D-G, Liu D-C, Zhang A-M, Xu H-X, Li B (2012) Mapping QTLs with epistatic effects and QTL × treatment interactions for salt tolerance at seedling stage of wheat. Euphytica 186:233–245CrossRefGoogle Scholar
  59. Yang J, Zhu J (2005) Methods for predicting superior genotypes under multiple environments based on QTL effects. Theor Appl Genet 110:1268–1274CrossRefGoogle Scholar
  60. Yang J, Hu C, Ye X, Zhu J (2005) QTLNetwork 2.0. Institute of Bioinformatics, Zhejiang University, HangzhouGoogle Scholar
  61. Yang J, Zhu J, Williams RW (2007) Mapping the genetic architecture of complex traits in experimental populations. Bioinformatics 23:1527–1536CrossRefGoogle Scholar
  62. Yang J, Hu C, Hu H, Yu R, Xia Z, Ye X, Zhu J (2008) QTLNetwork: mapping and visualizing genetic architecture of complex traits in experimental populations. Bioinformatics 24:721–723CrossRefGoogle Scholar
  63. Ziegler-Jöns A (1989) Gas-exchange of ears of cereals in response to carbon dioxide and light. Planta 178:164–175CrossRefGoogle Scholar

Copyright information

© The Genetics Society of Korea and Springer Nature B.V. 2018

Authors and Affiliations

  • Bahram Masoudi
    • 1
    Email author
  • Mohsen Mardi
    • 2
  • Eslam Majidi Hervan
    • 3
  • Mohammad Reza Bihamta
    • 4
  • Mohammad Reza Naghavi
    • 4
  • Babak Nakhoda
    • 3
  • Behnam Bakhshi
    • 5
  • Mehrzad Ahmadi
    • 1
  • Mohammad Taghi Tabatabaei
    • 6
  • Mohamad Hossein Dehghani Firouzabadi
    • 6
  1. 1.Seed and Plant Improvement InstituteAgricultural Research, Education and Extension Organization (AREEO)KarajIran
  2. 2.Department of GenomicsAgricultural Biotechnology Research Institute of Iran (ABRII)KarajIran
  3. 3.Department of Molecular PhysiologyAgricultural Biotechnology Research Institute of Iran (ABRII)KarajIran
  4. 4.Department of Plant BreedingThe University of TehranKarajIran
  5. 5.Horticulture Crops Research Department, Sistan Agricultural and Natural Resources Research and Education CenterAgricultural Research, Education and Extension Organization (AREEO)ZabolIran
  6. 6.Experimental Center of Agriculture and Natural ResourcesAgricultural Research, Education and Extension Organization (AREEO)YazdIran

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